We studied the breeding biology of Imperial Cormorant <i>Phalacrocorax atriceps</i> and Rock Shag <i>Phalacrocorax magellanicus</i> at Golfo San Jorge, Argentina, from 1991 to 1993. Dates in which first eggs were observed varied among seasons, being between mid October and mid November for Imperial Cormorants, and between late October and early December for Rock Shags. During 1993, 87% and 86% Imperial Cormorant and Rock Shag pairs, respectively, completed their clutches within four weeks after egg-laying started. The number of eggs per nest recorded one month after the start of egg laying for both species was statistically different among years. The average number of eggs per nest was similar between Imperial Cormorants and Rock Shags during 1993 (2.4 vs. 2.1, respectively) but significantly different during 1991 (1.9 vs. 0.8). During two of the study seasons the volume of the third egg was significantly smaller than the first and second eggs. The average length of the incubation period for Imperial Cormorants was estimated at 27.9 days. Hatching success of Imperial Cormorants was 38.2% (1991) and 52.5% (1992). Chick weight increase and culmen growth showed maximum values during the second and third weeks after hatching, reaching asymptotic values at 60 days. Chick tarsus growth reached maximum values during the first week and asymptotic values at 30 days. During 1993 mean number of chicks fledged per nest was 1.13 for Imperial Cormorants and 0.83 for Rock Shags.

Diets of breeding Imperial Cormorants Phalacrocorax atriceps were studied at two breeding colonies, Islas Blancas and Isla Arce, located approximately 30 km apart in an area subject to increasing fishing pressure off Central Patagonia, Argentina. The goal was to assess differences between locations and diet variation among stages of the breeding cycle. Pellet casts (403 and 358 pellets per colony, respectively) were collected from November 2002 to February 2003. Analyses of the pellets revealed that Imperial Cormorants at Islas Blancas and Isla Arce fed on at least 25 and 23 prey types, respectively. Fish showed the highest frequency of occurrence at both colonies (> 70%), followed by crustaceans and molluscs. Of the fish prey, Merluccius hubbsi (2271%), Engranlis anchoita (1651%) and Raneya brasiliensis (548%) showed the highest frequencies of occurrence, depending on the colony and breeding stage. At Islas Blancas, the consumption of fish and crustaceans was similar among breeding stages (incubation, young chicks and old chicks), while it was significantly different at Isla Arce. Overall contribution by frequency of occurrence showed that M. hubbsi was the most frequent prey at Islas Blancas (58%), and E. anchoita and Pleoticus muelleri were more frequent at Isla Arce (48 and 45%, respectively). Also, overall contribution by mass of the main fish prey indicated differences between colonies. Given the commercial value of the main prey species, cormorant feeding requirements and spatial ecological needs should be included as considerations in coastal fisheries management and future development.

Energy management during the breeding season is crucial for central place foragers since parents need to feed themselves and their offspring while being spatially and temporally constrained. In this work, we used overall dynamic body acceleration as a measure of activity and also to allude to the foraging energy expenditure of breeding Imperial cormorants Phalacrocorax atriceps. We also analyzed how changes in the time or energy allocated to different activities affected the foraging trip energy expenditure and estimated the daily food requirements of the species. Birds spent 42 % of the total energy flying to and from the feeding areas and 16 % floating at sea. The level of activity underwater was almost 1.5 times higher for females than for males. The most expensive diving phase in terms of rate of energy expenditure was descending though the water column. The total foraging trip energy expenditure was particularly sensitive to variation in the amount of time spent flying. During the breeding season, adult cormorants breeding along the Patagonian coast would consume approximately 10,000 tons of food.

Several studies have applied induced regurgitations to characterize the diet of cormorants, but none have presented quantitative information indicating complete stomach contents were obtained. Our goal was to test the value of induced regurgitations for the assessment and monitoring of Imperial Cormorant (Phalacrocorax atriceps) diet. Stomach samples were obtained from male and female breeding adults bringing food back to the colony during the chick rearing stage (n = 22) at Isla Arce, Argentina. Samples were obtained through induced regurgitation, and immediately afterward each individual was flushed with sea water. The diet of the Imperial Cormorant consisted of at least 23 prey taxa, mostly fish complemented by crustaceans, cephalopods and polychaetes. However, only Argentine anchovy (Engraulis anchoita) and rockcods (Patagonotothen spp.) showed a significant contribution by mass (70.7% and 25.3%, respectively). Analysis of similarity indicated that prey composition between samples obtained by induced regurgitation and those obtained by combining regurgitation followed by stomach flushing were similar in both the numerical frequency of all prey taxa recorded and the contribution by mass of the main prey. Our results show that induced regurgitation provides complete stomach contents, and thus validates the use of this technique for quantifying Imperial Cormorant diet composition.

Intersexual differences in the foraging behaviour have been examined in several seabird species, especially those exhibiting sexual size dimorphism. We studied intersex behavioural differences in the Imperial Cormorant (Phalacrocorax atriceps), a size dimorphic seabird. Twenty adults (11 females and 9 males), breeding at Punta León (43°04′S; 64°2′W), Chubut, Argentina, were equipped with loggers to measure tri-axial acceleration and depth, to calculate the foraging trip time/activity budgets and diving behaviour. Both sexes had similar foraging trip durations, spending a similar proportion of the foraging time flying and floating on the sea surface. However, females spent more time underwater, executing more and shallower dives. Females also recovered more quickly than males from dives performed to depths of less than 30 m and spent more time foraging along the bottom at any depth than males. We conclude that if allometric effects affect the foraging behaviour of Imperial Cormorants, they only do so during diving because no differences were observed in the total amount of time sexes spent flying or foraging.

Plasma biochemistries provide a complementary method for assessing physiological and nutritional status of free-ranging wild birds. Triglycerides, total protein and alkaline phosphatase were determined in 110 free-living Imperial Cormorant (Phalacrocorax atriceps) chicks aged 16-35 days, at Punta León (Argentina) during 2010 and 2011. Body mass at 30 days of age ("pre-fledging body condition", 2010 only) and body mass corrected by tarsus length at the time of blood sampling ("current body condition", 2011 only) were also determined. Variability of parameters by sex, hatching order, survival, age and breeding season was assessed, and the relationship between biochemical and morphometric indices was also explored. Morphometric indices were higher in A-chicks (pre-fledging body condition also varied with sex), and explained 35-55% of B-chick survival. Biochemistries differed significantly between breeding seasons, being higher in 2011. Alkaline phosphatase increased with age, and total protein was higher in A-chicks. Triglycerides and total protein accounted for 26% and 30%, respectively, of variation in current body condition; however, they did not forecast pre-fledging body condition. Lastly, total protein levels predicted B-chick survival (higher levels in surviving B-chicks), but their prognostic value was relatively low. The results suggest that unlike morphometric indices, the biochemistries chosen are valuable to assess individual body condition at the time of sampling, yet their applicability for predicting chick survival requires further evaluation.

Behavior by a foraging seabird during the breeding season can be examined by analyzing time invested throughout the foraging route to determine the presence of Area-Restricted Search (ARS) as well as other characteristics related to the shape of the foraging path and activity (flying, resting or diving). Forty-six Imperial Cormorants (Phalacrocorax atriceps), 18 males and 28 females breeding at Punta León, Argentina (43°04´S; 64°2´W), were fitted with GPS loggers recording one foraging trip (sampling interval: 1 second) in 2004, 2005, 2007 and 2008. Trip duration was longer, on average, for females (6.3 ± 1.9 h) than males (5.3 ± 1.6 h) (Mann-Whitney U test z 1,45 = 2.23, P = 0.026), but year had no significant effect on any of the path characteristics. ARS was detected in forty-three individuals, twenty-two of which made smaller scaled searches nested within larger ARS areas. Search scale was not correlated to maximum distance reached and did not differ between sexes nor years. Cluster analysis separated four types of behavior: short direct return trips (N = 7), long direct return trips (N = 31), random flight searches (N = 6) and loops (N = 2), with each behavioral type present in both sexes. Behavioral variability within a population may be due to differences in targeted prey type and spatio-temporal stability during the season, as well as to individual physiological constraints and life-history traits linked to how individuals search for food at sea.

The time seabirds have to forage is restricted while breeding, as time at sea must be balanced against the need to take turns with the partner protecting the nest site or offspring, and timing constraints change once the breeding season is over. Combined geolocator-immersion devices were deployed on eleven Imperial Shags (four males and seven females) in Argentina (43°04′S; 64°2′W) in November 2006 and recovered in November 2007. During the breeding season, females foraged throughout the morning, males exclusively in the afternoon, and variability between individuals was low. Outside the breeding season, both sexes foraged throughout the day, and variability between individuals was high. Timing differences may be explained by higher constraints on foraging or greater demands of parental duties experienced by the smaller sex, females in this case. Sexual differences in reproductive role, feeding habits or proficiency can also lead to segregation in timing of foraging, particularly while breeding.

Understanding the dynamics and causes of population trends are essential for seabird conservation. Long-term studies of seabirds at high-latitude (Antarctic, sub-Antarctic and Arctic) regions have shown contrasting species-specific trends in population size in response to climate change and anthropogenic pressures. We have studied for the last 20 years (1992?2012) the population trends of seven seabird species that breed in the Beagle Channel, south-eastern Tierra del Fuego and at Staten Island, a sub-Antarctic region in southern Argentina. The numbers of Magellanic and Gentoo Penguins increased significantly since 1992 (by[15 % year-1). In comparison, the populations of Imperial Cormorants, Dolphin Gulls and Kelp Gulls increased at slower rates ( % year-1), while the Rock Cormorant population even decreased by 1.3 % year-1. At Staten Island, the numbers of Rockhopper Penguins decreased by 24 % between the censuses of 1998 and 2010, whereas the population of Magellanic Penguins increased by 227 % during the same period. Over the study period, air and sea-surface temperatures remained stable in our study area, suggesting that the detected population changes are not driven by the climate. This finding contrasts with the detected links between increasing temperature trends and seabird population changes reported from Antarctic and Arctic regions. The level of tourism and size of the permanent human population has increased in the Beagle Channel area during the last 20 years and could be responsible for the increase of gull populations. The seabird species that received the highest number of visitors (Imperial Cormorants and penguin species) seem to be adapted or at least indifferent to pressures exerted by tourism, as their populations increased during the study period. In addition, increasing numbers of seabirds in the area may generally be leading to higher abundances of scavenging species (e.g. gulls).

Air-breathing animals diving to forage can optimize time underwater by diving with just enough oxygen for the projected performance underwater. By so doing they surface with minimal body oxygen levels, which leads to maximal rates of oxygen uptake. We examined whether imperial cormorants Phalacrocorax atriceps adhere to this by examining dive: pause ratios in birds diving for extended, continuous periods to constant depths, assuming that the oxygen used underwater was exactly replenished by the periods at the surface. Examination of the cumulative time spent in surface pauses relative to the cumulative time spent in diving showed that surface pauses increase according to a power curve function of time spent in the dive or water depth. In a simplistic model we considered the rate at which birds expended energy underwater to be constant and that the rate of oxygen replenishment during the surface pause was directly proportional to the oxygen deficit. We then worked out values for the rate constant for the surface pause before using this constant to examine bird body oxygen levels immediately pre- and post dive. The model predicted that imperial cormorants do not submerge with just enough oxygen to cover their projected dive performance but rather dive with substantial reserves, although these reserves decrease with increasing dive depth/duration. We speculate that these oxygen reserves may be used to enhance bird survival when rare events, such as the appearance of predators or discovery of large prey requiring extended handling time, occur. The form of the oxygen saturation curve over time at the surface means that the time costs for maintaining constant oxygen reserves become particularly onerous for long, deep dives, so the observed decrease in reserves with increasing dive duration is expected in animals benefiting by optimizing for time